Interactions between beech and oak seedlings can modify the effects of hotter droughts and the onset of hydraulic failure.

Mixing species with contrasting resource use strategies could reduce forest vulnerability to extreme events. Yet, how species diversity affects seedling hydraulic responses to heat and drought, including mortality risk, is largely unknown. Using open-top chambers, we assessed how, over several years, species interactions (monocultures vs mixtures) modulate heat and drought impacts on the hydraulic traits of juvenile European beech and pubescent oak. Using modeling, we estimated species interaction effects on timing to drought-induced mortality and the underlying mechanisms driving these impacts. We show that mixtures mitigate adverse heat and drought impacts for oak (less negative leaf water potential, higher stomatal conductance, and delayed stomatal closure) but enhance them for beech (lower water potential and stomatal conductance, narrower leaf safety margins, faster tree mortality). Potential underlying mechanisms include oak's larger canopy and higher transpiration, allowing for quicker exhaustion of soil water in mixtures. Our findings highlight that diversity has the potential to alter the effects of extreme events, which would ensure that some species persist even if others remain sensitive. Among the many processes driving diversity effects, differences in canopy size and transpiration associated with the stomatal regulation strategy seem the primary mechanisms driving mortality vulnerability in mixed seedling plantations.

Temperature sensing: A potassium channel as cold sensor in the rain tree Samanea saman.

The rain tree Samanea saman folds its leaves upon rainfall. New results now indicate that rain perception is in fact a temperature-sensing process, and that Samanea possess an ion channel with a strong temperature sensitivity that is involved in leaf movement.

The Surprising Dynamics of Electrochemical Coupling at Membrane Sandwiches in Plants.

Transport processes across membranes play central roles in any biological system. They are essential for homeostasis, cell nutrition, and signaling. Fluxes across membranes are governed by fundamental thermodynamic rules and are influenced by electrical potentials and concentration gradients. Transmembrane transport processes have been largely studied on single membranes. However, several important cellular or subcellular structures consist of two closely spaced membranes that form a membrane sandwich. Such a dual membrane structure results in remarkable properties for the transport processes that are not present in isolated membranes. At the core of membrane sandwich properties, a small intermembrane volume is responsible for efficient coupling between the transport systems at the two otherwise independent membranes. Here, we present the physicochemical principles of transport coupling at two adjacent membranes and illustrate this concept with three examples. In the supplementary material, we provide animated PowerPoint presentations that visualize the relationships. They could be used for teaching purposes, as has already been completed successfully at the University of Talca.

TPC1-Type Channels in Physcomitrium patens: Interaction between EF-Hands and Ca2.

Two-pore channels (TPCs) are members of the superfamily of ligand-gated and voltage-sensitive ion channels in the membranes of intracellular organelles of eukaryotic cells. The evolution of ordinary plant TPC1 essentially followed a very conservative pattern, with no changes in the characteristic structural footprints of these channels, such as the cytosolic and luminal regions involved in Ca2+ sensing. In contrast, the genomes of mosses and liverworts encode also TPC1-like channels with larger variations at these sites (TPC1b channels). In the genome of the model plant Physcomitrium patens we identified nine non-redundant sequences belonging to the TPC1 channel family, two ordinary TPC1-type, and seven TPC1b-type channels. The latter show variations in critical amino acids in their EF-hands essential for Ca2+ sensing. To investigate the impact of these differences between TPC1 and TPC1b channels, we generated structural models of the EF-hands of PpTPC1 and PpTPC1b channels. These models were used in molecular dynamics simulations to determine the frequency with which calcium ions were present in a coordination site and also to estimate the average distance of the ions from the center of this site. Our analyses indicate that the EF-hand domains of PpTPC1b-type channels have a lower capacity to coordinate calcium ions compared with those of common TPC1-like channels.

Global trends in phenotypic plasticity of plants.

Predicting plastic responses is crucial to assess plant species potential to adapt to climate change, but little is known about which factors drive the biogeographical patterns of phenotypic plasticity in plants. Theory predicts that climatic variability would select for increased phenotypic plasticity, whereas evidence indicates that stressful conditions can limit phenotypic plasticity. Using a meta-analytic, phylogeny-corrected approach to global data on plant phenotypic plasticity, we tested whether latitude, climate, climatic variability and/or stressful conditions are predictors of plastic responses at a biogeographical scale. We found support for a positive association between phenotypic plasticity and climatic variability only for plasticity in allocation. Plasticity in leaf morphology, size and physiology were positively associated with mean annual temperature. We also found evidence that phenotypic plasticity in physiology is limited by cold stress. Overall, plant plastic responses to non-climatic factors were stronger than responses to climatic factors. However, while climatic conditions were associated with plant plastic responses to climatic factors, they generally did not relate to plastic responses to other abiotic or biotic factors. Our study highlights the need to consider those factors that favour and limit phenotypic plasticity in order to improve predictive frameworks addressing plant species' potential to adapt to climate change.

Future paths for the 'exploitative segregation of plant roots' model.

The exploitative segregation of plant roots (ESPR) is a theory that uses a game-theoretical model to predict plant root foraging behavior in space. The original model returns the optimal root distribution assuming exploitative competition between a pair of identical plants in soils with homogeneous resource dynamics. In this short communication, we explore avenues to develop this model further. We discuss: (i) the response of single plants to soil heterogeneity; (ii) the variability of the plant response under uneven competition scenarios; (iii) the importance of accounting for the constraints and limitations to root growth that may be imposed from the plant shoot; (iv) the importance of root functional traits to predict root foraging behavior; (v) potential model extensions to investigate facilitation by incorporating facilitative traits to roots, and (vi) the possibility of allowing plants to tune their response by accounting for non-self and non-kin root recognition. For each case, we introduce the topic briefly and present possible ways to encode those ingredients in the mathematical equations of the ESPR model, providing preliminary results when possible.

The exploitative segregation of plant roots.

Plant roots determine carbon uptake, survivorship, and agricultural yield and represent a large proportion of the world's vegetation carbon pool. Study of belowground competition, unlike aboveground shoot competition, is hampered by our inability to observe roots. We developed a consumer-resource model based in game theory that predicts the root density spatial distribution of individual plants and tested the model predictions in a greenhouse experiment. Plants in the experiment reacted to neighbors as predicted by the model's evolutionary stable equilibrium, by both overinvesting in nearby roots and reducing their root foraging range. We thereby provide a theoretical foundation for belowground allocation of carbon by vegetation that reconciles seemingly contradictory experimental results such as root segregation and the tragedy of the commons in plant roots.

Importance of overstorey attributes for understorey litter production and nutrient cycling in European forests.

BACKGROUND: In contrast with the negligible contribution of the forest understorey to the total aboveground phytobiomass of a forest, its share in annual litter production and nutrient cycling may be more important. Whether and how this functional role of the understorey differs across forest types and depends upon overstorey characteristics remains to be investigated. METHODS: We sampled 209 plots of the FunDivEUROPE Exploratory Platform, a network of study plots covering local gradients of tree diversity spread over six contrasting forest types in Europe. To estimate the relative contribution of the understorey to carbon and nutrient cycling, we sampled non-lignified aboveground understorey biomass and overstorey leaf litterfall in all plots. Understorey samples were analysed for C, N and P concentrations, overstorey leaf litterfall for C and N concentrations. We additionally quantified a set of overstorey attributes, including species richness, proportion of evergreen species, light availability (representing crown density) and litter quality, and investigated whether they drive the understorey's contribution to carbon and nutrient cycling. RESULTS AND CONCLUSIONS: Overstorey litter production and nutrient stocks in litterfall clearly exceeded the contribution of the understorey for all forest types, and the share of the understorey was higher in forests at the extremes of the climatic gradient. In most of the investigated forest types, it was mainly light availability that determined the contribution of the understorey to yearly carbon and nutrient cycling. Overstorey species richness did not affect the contribution of the understorey to carbon and nutrient cycling in any of the investigated forest types.

UV radiation increases phenolic compound protection but decreases reproduction in Silene littorea.

Plants respond to changes in ultraviolet (UV) radiation both morphologically and physiologically. Among the variety of plant UV-responses, the synthesis of UV-absorbing flavonoids constitutes an effective non-enzymatic mechanism to mitigate photoinhibitory and photooxidative damage caused by UV stress, either reducing the penetration of incident UV radiation or acting as quenchers of reactive oxygen species (ROS). In this study, we designed a UV-exclusion experiment to investigate the effects of UV radiation in Silene littorea. We spectrophotometrically quantified concentrations of both anthocyanins and UV-absorbing phenolic compounds in petals, calyces, leaves and stems. Furthermore, we analyzed the UV effect on the photosynthetic activity in hours of maximum solar radiation and we tested the impact of UV radiation on male and female reproductive performance. We found that anthocyanin concentrations showed a significant decrease of about 20% with UV-exclusion in petals and stems, and a 30% decrease in calyces. The concentrations of UV-absorbing compounds under UV-exclusion decreased by approximately 25% in calyces and stems, and 12% in leaves. Photochemical efficiency of plants grown under UV decreased at maximum light stress, reaching an inhibition of 58% of photosynthetic activity, but their ability to recover after light-stress was not affected. In addition, exposure to UV radiation did not affect ovule production or seed set per flower, but decreased pollen production and total seed production per plant by 31% and 69%, respectively. Our results demonstrate that UV exposure produced opposing effects on the accumulation of plant phenolic compounds and reproduction. UV radiation increased the concentration of phenolic compounds, suggesting a photoprotective role of plant phenolics against UV light, yet overall reproduction was compromised.

Author Correction: The GenTree Dendroecological Collection, tree-ring and wood density data from seven tree species across Europe.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Intraspecific perspective of phenotypic coordination of functional traits in Scots pine.

Functional traits have emerged as a key to understand species responses to environmental conditions. The concerted expression of multiple traits gives rise to the phenotype of each individual, which is the one interacting with the environment and evolving. However, patterns of trait covariation and how they vary in response to environmental conditions remain poorly understood, particularly at the intraspecific scale. Here, we have measured traits at different scales and in different organs, and analysed their covariation in a large number of conspecifics distributed in two contrasting environments. We expected significant correlations among traits, not only within clusters of traits as found in global, multispecies studies, but also among clusters, with more relationships within clusters, due to genetic constraints, and among clusters due to more coordinated phenotypes than community level, multispecies studies. We surveyed 100 Pinus sylvestris trees in a Mediterranean mountainous area distributed in two contrasting elevations. We measured 13 functional traits, in three clusters (leaf, stem and whole-plant traits), and analysed their variation and coordination. We found significant coordination among traits belonging to different clusters that reveals coordinated phenotypes. However, we found fewer correlations within trait clusters than initially expected. Trait correlation structures (number, intensity and type of correlations among traits) differed among individuals at different elevations. We observed more correlations within trait clusters at low elevation compared to those at high elevation. Moreover, the higher number of correlations among different trait clusters and the lower trait variation at the higher elevation suggests that variability decreases under more stressful conditions. Altogether, our results reveal that traits at intraspecific scale are coordinated in a broad network and not only within clusters of traits but also that this trait covariation is significantly affected by environmental conditions.

The GenTree Dendroecological Collection, tree-ring and wood density data from seven tree species across Europe.

The dataset presented here was collected by the GenTree project (EU-Horizon 2020), which aims to improve the use of forest genetic resources across Europe by better understanding how trees adapt to their local environment. This dataset of individual tree-core characteristics including ring-width series and whole-core wood density was collected for seven ecologically and economically important European tree species: silver birch (Betula pendula), European beech (Fagus sylvatica), Norway spruce (Picea abies), European black poplar (Populus nigra), maritime pine (Pinus pinaster), Scots pine (Pinus sylvestris), and sessile oak (Quercus petraea). Tree-ring width measurements were obtained from 3600 trees in 142 populations and whole-core wood density was measured for 3098 trees in 125 populations. This dataset covers most of the geographical and climatic range occupied by the selected species. The potential use of it will be highly valuable for assessing ecological and evolutionary responses to environmental conditions as well as for model development and parameterization, to predict adaptability under climate change scenarios.

Soil fertility and rainfall during specific phenological phases affect seed trait variation in a widely distributed Neotropical tree, Copaifera langsdorffii.

PREMISE: Knowledge of intra-specific variation in seed traits and its environmental determinants is important for predicting plant responses to environmental changes. Here, we tested the hypothesis that differences in soil fertility and rainfall during specific phenological phases drive variation in seed traits in a widely distributed tree, Copaifera langsdorffii. We also tested the hypothesis that climatic heterogeneity increases within-plant variation in seed traits. METHODS: Inter- and intra-population and within-plant variation in seed mass, number, and seed size/seed number were evaluated for 50 individuals from five populations distributed along a rainfall gradient and occurring on varying soil types. Using multivariate approaches, we tested the effects of soil fertility characteristics and rainfall in five reproductive phenological phases on seed traits. RESULTS: The seed traits varied greatly both among populations and within plants. Inter-population variation in seed mass was driven by total rainfall during fruit development, and variation in seed number was influenced by total rainfall during the dry season before the reproductive phase. Phosphorus levels and potential acidity of the soil also explained the variations in seed mass and seed mass/seed number, respectively. A positive association between intra-annual variation in rainfall and within-plant variation in seed mass and seed number was found. CONCLUSION: Both rainfall during specific reproductive phases and soil conditions shape the variation in the seed mass and number of C. langsdorffii. Environment-driven seed trait variation may contribute to this species' broad niche breadth, which in turn may determine the species' persistence under future climatic conditions.

Responses of Aspen Leaves to Heatflecks: Both Damaging and Non-Damaging Rapid Temperature Excursions Reduce Photosynthesis.

During exposure to direct sunlight, leaf temperature increases rapidly and can reach values well above air temperature in temperate forest understories, especially when transpiration is limited due to drought stress, but the physiological effects of such high-temperature events are imperfectly understood. To gain insight into leaf temperature changes in the field and the effects of temperature variation on plant photosynthetic processes, we studied leaf temperature dynamics under field conditions in European aspen (Populus tremula L.) and under nursery conditions in hybrid aspen (P. tremula × P. tremuloides Michaux), and further investigated the heat response of photosynthetic activity in hybrid aspen leaves under laboratory conditions. To simulate the complex fluctuating temperature environment in the field, intact, attached leaves were subjected to short temperature increases ("heat pulses") of varying duration over the temperature range of 30 °C-53 °C either under constant light intensity or by simultaneously raising the light intensity from 600 µmol m-2 s-1 to 1000 µmol m-2 s-1 during the heat pulse. On a warm summer day, leaf temperatures of up to 44 °C were measured in aspen leaves growing in the hemiboreal climate of Estonia. Laboratory experiments demonstrated that a moderate heat pulse of 2 min and up to 44 °C resulted in a reversible decrease of photosynthesis. The decrease in photosynthesis resulted from a combination of suppression of photosynthesis directly caused by the heat pulse and a further decrease, for a time period of 10-40 min after the heat pulse, caused by subsequent transient stomatal closure and delayed recovery of photosystem II (PSII) quantum yield. Longer and hotter heat pulses resulted in sustained inhibition of photosynthesis, primarily due to reduced PSII activity. However, cellular damage as indicated by increased membrane conductivity was not found below 50 °C. These data demonstrate that aspen is remarkably resistant to short-term heat pulses that are frequent under strongly fluctuating light regimes. Although the heat pulses did not result in cellular damage, heatflecks can significantly reduce the whole plant carbon gain in the field due to the delayed photosynthetic recovery after the heat pulse.

Phenotypic plasticity closely linked to climate at origin and resulting in increased mortality under warming and frost stress in a common grass.

Phenotypic plasticity is important for species responses to global change and species coexistence. Phenotypic plasticity differs among species and traits and changes across environments. Here, we investigated phenotypic plasticity of the widespread grass Arrhenatherum elatius in response to winter warming and frost stress by comparing phenotypic plasticity of 11 geographically and environmentally distinct populations of this species to phenotypic plasticity of populations of different species originating from a single environment. The variation in phenotypic plasticity was similar for populations of a single species from different locations compared to populations of functionally and taxonomically diverse species from one environment for the studied traits (leaf biomass production and root integrity after frost) across three indices of phenotypic plasticity (RDPI, PIN, slope of reaction norm). Phenotypic plasticity was not associated with neutral genetic diversity but closely linked to the climate of the populations' origin. Populations originating from warmer and more variable climates showed higher phenotypic plasticity. This indicates that phenotypic plasticity can itself be considered as a trait subject to local adaptation to climate. Finally, our data emphasize that high phenotypic plasticity is not per se positive for adaptation to climate change, as differences in stress responses are resulting in high phenotypic plasticity as expressed by common plasticity indices, which is likely to be related to increased mortality under stress in more plastic populations.

Selfing and Drought-Stress Strategies Under Water Deficit for Two Herbaceous Species in the South American Andes.

Angiosperms are highly diverse in their reproductive systems, including predominantly selfing, exclusive outcrossing, and mixed mating systems. Even though selfing can have negative consequences on natural populations, it has been proposed that plants having a predominantly selfing strategy are also associated with fast development strategies through time limitation mechanisms that allow them to complete their life cycle before the onset of severe drought. This relationship might be affected by the challenges imposed by global change, such as a decrease in pollinator availability and the earlier and more severe onset of droughts. In this work, our aim was to investigate whether selfing is correlated with a dehydration avoidance strategy, and how this could affect drought resistance and survival in two species with different types of selfing: pollinator-independent delayed selfing (Schizanthus grahamii) and pollinator-dependent selfing (Schizanthus hookeri), representing a gradient in selfing rates. We hypothesize that delayed selfing species and highly selfing populations will show "fast" plant traits whereas we will find no pattern in more outcrossed populations of the pollinator-dependent species. However, we predicted that high selfing populations would have lower survival rates when exposed to chronic drought early in their development since fast traits imply physiological compromises that will affect their drought survival. To evaluate these hypotheses, we characterized different physiological and morphological traits in response to two contrasting treatments (moist and dry) in a total of six populations of the two species. We found a relationship between the delayed selfing species and a dehydration avoidance strategy and also with low drought survival. Our work offers evidence to support the importance of abiotic factors, such as drought, on the possible variation in selfing rates on natural populations, and the effect that this mating system could have in their ability to face new environmental conditions such as those imposed by climate change.

Increased root investment can explain the higher survival of seedlings of 'mesic' Quercus suber than 'xeric' Quercus ilex in sandy soils during a summer drought.

In Mediterranean-type ecosystems, drought is considered the main ecological filter for seedling establishment. The evergreen oaks Quercus ilex L. and Quercus suber L. are two of the most abundant tree species in the Mediterranean Basin. Despite their shared evergreen leaf habit and ability to resist low soil water potentials, traditionally it has been suggested that Q. ilex is better suited to resist dry conditions than Q. suber. In this study, we examined how seedlings of Q. ilex and Q. suber grown in sandy soils responded to different levels of water availability using natural dry conditions and supplemental watering. Specifically, we estimated survival and water status of seedlings and explored the role of acorn mass and belowground biomass in seedling performance. To our surprise, Q. suber was better able to survive the summer drought in our experiment than Q. ilex. Nearly 55% of the Q. suber seedlings remained alive after a 2-month period without rain or supplemental water, which represents almost 20% higher survival than Q. ilex over the same period. At the end of the dry period, the surviving seedlings of Q. suber had strikingly higher water potential, potential maximum quantum yield of photosystem II (Fv/Fm) and stomatal conductance (gs) than those of Q. ilex. Acorn mass was associated with the probability of survival under dry conditions; however, it did not explain the differences in survival or water status between the species. In contrast, Q. suber had a higher root ratio and root:shoot ratio than Q. ilex and these traits were positively associated with predawn leaf water potential, Fv/Fm, gs and survival. Taken together, our results suggest that the higher relative investment in roots by Q. suber when growing in a sandy acidic substrate allowed this species to maintain better physiological status and overall condition than Q. ilex, increasing its probability of survival in dry conditions.

Climatic factors shaping intraspecific leaf trait variation of a neotropical tree along a rainfall gradient.

Intraspecific trait variation has been singled out as an important mechanism by which individuals can cope with environmental variations and avoid local extinctions. Here we evaluate variation in metamer traits (i.e., traits associated with internodes, petioles and their corresponding leaves) and parameters of chlorophyll fluorescence within and among populations of a neotropical tree, Copaifera langsdorffii. We also evaluated phenotypic plasticity in natural settings comparing traits between shade and sun-exposed metamers. We selected six populations along a climatic gradient ranging from semi-arid to humid and representing three different biomes (Caatinga, Cerrado, and Atlantic Forest). Local climatic conditions significantly affected the morphological and physiological traits of populations. Trait variation among populations was explained mainly by aridity index and evapotranspiration. Individuals from drier regions had lower specific leaf area (SLA), lower investment in leaf area per total dry mass of metamer (LARm), lower specific petiole length (SPL) and lower potential quantum yield (Fv/Fm, only for sun-exposed metamers). Populations from locations with greater environmental heterogeneity (interannual variation) had greater plasticity in response to light for Fv/Fm and electron transport rate (ETR) and morphological traits related to the hydraulic and biomechanical aspects of the leaves (petiole length, internode length and SPL). High intraspecific variation in metamer traits in C. langsdorffii coupled with its ability to modify these traits in response to different climate conditions can explain the success of the species over a range of different habitats and represent important factors for the persistence of this species in the face of climate change.

Habitat fragmentation is linked to cascading effects on soil functioning and CO2 emissions in Mediterranean holm-oak-forests.

We studied key mechanisms and drivers of soil functioning by analyzing soil respiration and enzymatic activity in Mediterranean holm oak forest fragments with different influence of the agricultural matrix. For this, structural equation models (SEM) were built including data on soil abiotic (moisture, temperature, organic matter, pH, nutrients), biotic (microbial biomass, bacterial and fungal richness), and tree-structure-related (basal area) as explanatory variables of soil enzymatic activity and respiration. Our results show that increased tree growth induced by forest fragmentation in scenarios of high agricultural matrix influence triggered a cascade of causal-effect relations, affecting soil functioning. On the one hand, soil enzymatic activity was strongly stimulated by the abiotic (changes in pH and microclimate) and biotic (microbial biomass) modifications of the soil environment arising from the increased tree size and subsequent soil organic matter accumulation. Soil CO2 emissions (soil respiration), which integrate releases from all the biological activity occurring in soils (autotrophic and heterotrophic components), were mainly affected by the abiotic (moisture, temperature) modifications of the soil environment caused by trees. These results, therefore, suggest that the increasing fragmentation of forests may profoundly impact the functioning of the plant-soil-microbial system, with important effects over soil CO2 emissions and nutrient cycling at the ecosystem level. Forest fragmentation is thus revealed as a key albeit neglected factor for accurate estimations of soil carbon dynamics under global change scenarios.